遮荫对弗吉尼亚栎苗木生长及光合-荧光参数的影响
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摘要
[目的]揭示美国引进树种弗吉尼亚栎幼苗对光照强度的响应特征,为弗吉尼亚栎的引种驯化和高效栽培提供科学依据。[方法]采用人工遮荫法,研究3种光环境下(全光——100%自然光、中度遮荫——50%自然光和强度遮荫——25%自然光)弗吉尼亚栎苗木生长、光合作用以及叶绿素荧光参数的变化。[结果]表明:中度遮荫明显促进弗吉尼亚栎苗高生长(P<0.05),但对地径生长影响不明显;而强度遮荫条件下,地径生长受到明显抑制(P<0.05),苗高虽有所下降,但与对照相比差异不明显。对弗吉尼亚栎光响应曲线的观测表明:与全光条件相比,中度遮荫条件下,弗吉尼亚栎叶片暗呼吸速率(Rd)、光饱和点(LSP)以及光补偿点(LCP)均明显下降(P<0.05),说明适度的遮荫有助于提高弗吉尼亚栎对光的利用效率,促进干物质的积累;而强度遮荫条件下,光补偿点和光饱和点明显上升,暗呼吸速率明显增加(P<0.05)。遮荫并未对表观量子效率(α)和最大净光合速率(P_(max))产生明显的影响。进一步对叶绿素荧光参数分析发现,遮荫条件下,PSⅡ最大光化学量子产量(F_v/F_m)和实际光量子产量(Yield)有所增加,但均在正常范围,说明遮荫对PSⅡ活性并未产生胁迫。[结论]弗吉尼亚栎幼苗对遮荫均具有一定的耐受性,中度遮荫有利于苗高生长;而强度遮荫不利于干物质的积累。因此,建议在营林以及人工栽培过程中维持50%以上的自然光照。
[Objective] To discover the response to light intensity of Quercus virginiana, and the effects of different levels of shading on growth, photosynthesis and chlorophyll fluorescence parameters. [Method] Three light levels(full light-100% natural light, moderate shading-50% natural light and high shading-25% natural light) were set in greenhouse by covering different layers of shading net. The growth, chlorophyll content index, photosynthesis and chlorophyll fluorescence parameters were determined during the experiment. [Result] The results showed that the height growth of seedlings increased significantly(P<0.05) under moderate shading condition, while no obvious change was observed in the growth of basal diameter. Under high shading conditions, the height growth changed a little compared with control, whereas the growth of basal diameter was inhibited significantly(P<0.05). Under moderate shading, the dark respiration rate(R_d), light compensation point(LCP) and light saturation point(LSP) declined and whereas increased significantly under high shading, which showed that moderate shading might be helpful to improve the light use efficiency and increase the dry mass accumulation in leaves of Q. virginiana. There were no obvious changes in apparent quantum yield(α) and the maximum net photosynthetic rate(P_(max)). Further analysis of chlorophyll fluorescence parameters showed that PSII primary energy conversion efficiency(F_v/F_m) and photochemical efficiency of PSII in the light(Yield) remained a reasonable range under two light levels, which indicated that the PSII activity might be not damaged. [Conclusion] These results indicate that Q. virginiana could tolerate certain shading conditions, however, high shading was considered to be detrimental to the accumulation of dry matter, and light condition with more than 50% natural light would be recommended during the silviculture and nursery cultivation.
[Objective] To discover the response to light intensity of Quercus virginiana, and the effects of different levels of shading on growth, photosynthesis and chlorophyll fluorescence parameters. [Method] Three light levels(full light-100% natural light, moderate shading-50% natural light and high shading-25% natural light) were set in greenhouse by covering different layers of shading net. The growth, chlorophyll content index, photosynthesis and chlorophyll fluorescence parameters were determined during the experiment. [Result] The results showed that the height growth of seedlings increased significantly(P<0.05) under moderate shading condition, while no obvious change was observed in the growth of basal diameter. Under high shading conditions, the height growth changed a little compared with control, whereas the growth of basal diameter was inhibited significantly(P<0.05). Under moderate shading, the dark respiration rate(R_d), light compensation point(LCP) and light saturation point(LSP) declined and whereas increased significantly under high shading, which showed that moderate shading might be helpful to improve the light use efficiency and increase the dry mass accumulation in leaves of Q. virginiana. There were no obvious changes in apparent quantum yield(α) and the maximum net photosynthetic rate(P_(max)). Further analysis of chlorophyll fluorescence parameters showed that PSII primary energy conversion efficiency(F_v/F_m) and photochemical efficiency of PSII in the light(Yield) remained a reasonable range under two light levels, which indicated that the PSII activity might be not damaged. [Conclusion] These results indicate that Q. virginiana could tolerate certain shading conditions, however, high shading was considered to be detrimental to the accumulation of dry matter, and light condition with more than 50% natural light would be recommended during the silviculture and nursery cultivation.
引文
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[2] 刘建锋,杨文娟,江泽平.等.遮荫对濒危植物崖柏光合作用和叶绿素荧光参数的影响[J].生态学报,2011,31(20):5999-6004.
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[4] 李东胜,白庆红,李永杰,等.光照条件对蒙古栎幼苗生长特性和光合特征的影响[J].生态学杂志,2017,36(10):2744-2750.
[5] 贺顺钦,王发其.辽东栎苗木早期生长与光的关系[J].林业科学研究,2001,14(6):697-700.
[6] 陈秋夏,廖亮,郑坚,等.光照强度对青冈栎容器苗生长和生理特征的影响[J].林业科学,2011,47(12):53-58.
[7] 徐飞,郭卫华,徐伟红,等.不同光环境对麻栎和刺槐幼苗生长和光合特征的影响[J].生态学报,2010,30(12):3098-3107.
[8] Haller J M.Quercus virginiana:the southern live oak[J].Arbor Age,1992,12(5):30.
[9] Harms W R.Quercus virginiana Mill.live oak[M]// Burns R M,Honkala B H.Silvics of North America:Vol.2 hardwoods.Agric.handb.654.Washington,DC:U.S.Department of Agriculture,Forest Service,1990:751-754.
[10] 陈益泰,陈雨春,黄一青,等.抗风耐盐常绿树种弗吉尼亚栎引种初步研究[J].林业科学研究,2007,20(4):542-546.
[11] 陈益泰,孙海菁,王树凤,等.5种北美栎树在我国长三角地区的引种生长表现[J].林业科学研究,2013,26(3):344-351.
[12] Cavender-Bares J M,Kitajima K,Bazzaz F A.Multiple trait associations in relation to habitat differentiation among 17 oak species in North Central Florida[J].Ecological Monographs,2004,74(4):635-662.
[13] Spector T,Putz F E.Crown retreat of open-grown Southern live oaks (Quercus virginiana) due to canopy encroachment in Florida,USA[J].Forest Ecology and Management,2006,228(1-3):168-176.
[14] 王树凤,胡韵雪,李志兰,等.盐胁迫对弗吉尼亚栎生长及矿质离子吸收、运输和分配的影响[J].生态学报,2010,30(17):4609-4616.
[15] 王树凤,胡韵雪,孙海菁,等.盐胁迫对2种栎树苗期生长和根系生长发育的影响[J].生态学报,2014,34(4):1021-1029.
[16] 杨舒婷,曲博,李谦盛,等.弗吉尼亚栎幼苗对高温胁迫的生理响应[J].江西农业大学学报,2015,37(1):90-95.
[17] 宋以刚,邓敏,王妍婷.弗吉尼亚栎种子萌发特性[J].生态学杂志,2015,4(5):1295-1300.
[18] Ghasemi M,Arzani K,Yadollahi A,et al.Estimate of leaf chlorophyll and nitrogen content in asian pear (Pyrus serotina Rehd.) by CCM-200[J].Notulae Scientia Biologicae,2011,3(1):91-94.
[19] 叶子飘,于强.一个光合作用光响应新模型与传统模型的比较[J].沈阳农业大学学报,2007,38(6):771-775.
[20] 叶子飘,于强.光合作用光响应模型的比较[J].植物生态学报,2008,32(6):1356-1361.
[21] Maxwell K,Johnson G N.Chlorophyll fluorescence—A practical guide[J].Journal of Experimental Botany,2000,51(345):659-668.
[22] 张守仁.叶绿素荧光动力学参数的意义及讨论[J].植物学通报,1999,16(4):444-448.
[23] 吴飞燕,伊力塔,李修鹏,等.不同光照强度对石栎幼苗叶绿素含量及叶绿素荧光参数的影响[J].东北农业大学学报,2012,43(4):88-92.
[24] 潘瑞炽.植物生理学[M].北京:高等教育出版社,2008:61-62.
[25] 石玉波,卓丽环,辛雅芬,等.不同遮光处理对白栎光合生理特性的影响[J].安徽农业科学,2010,38(2):956-958.
[26] Dai Y J,Shen Z G,Liu Y,et al.Effects of shade treatments on the photosynthetic capacity,chlorophyll fluorescence,and chlorophyll content of Tetrastigma hemsleyanum Diels et Gilg[J].Environmental and Experimental Botany,2009,65(2-3):177-182.
[27] Falster D S,Westoby M.Tradeoffs between height growth rate,stem persistence and maximum height among plant species in a post-fire succession[J].Oikos,2005,111(1):57-66.
[28] Gardiner E S,Hodges J D.Growth and biomass distribution of cherrybark oak (Quercus pagoda Raf) seedlings as influenced by light availability[J].Forest Ecology and Management,1998,108(1-2):127-134.
[29] 闫兴富,曹敏.林窗对热带雨林冠层树种绒毛番龙眼幼苗生长的影响[J].应用生态学报,2008,19( 2):238-244.
[30] 杨兴洪,邹琦,赵世杰.遮荫和全光下生长的棉花光合作用和叶绿素荧光特征[J].植物生态学报,2005,29 (1):8-15
[31] 薛思雷,王庆成,孙欣欣,等.遮荫对水曲柳和蒙古栎光合、生长和生物量分配的影响[J].植物研究,2012,32(3):354-359.
[32] 郭志华,张旭东,黄玲玲,等.落叶阔叶树种蒙古栎(Quercus mongolica)对林缘不同光环境光能和水分的利用[J].生态学报,2006,26(4):1047-1056.
[33] Richardson A D,Berlyn G P.Spectral reflectance and photosynthetic properties of Betula papyrifera (Betulaceae) leaves along an elevational gradient on Mt.Mansfield,Vermont,USA[J].American Journal of Botany,2005,89(1):88-94.
[34] 梁文斌,聂东伶,吴思政,等.遮荫对短梗大参苗木光合作用及生长的影响[J].生态学杂志,2015,34(2):413-419.
[35] 唐敏,翟秀明,姚永红,等.不同茶树品种(系)叶绿素荧光参数季节性差异研究[J].西南农业学报,2015,28(1):79-83
[36] Yang H L,Yang X,Zhang Y G,et al.Chlorophyll fluorescence tracks seasonal variations of photosynthesis from leaf to canopy in a temperate forest[J].Global Change Biology,2017,23(7):2874-2886.
[37] Crous K Y,Zaragoza-Castells J,Ellsworth D S,et al.Light inhibition of leaf respiration in field-grown Eucalyptus saligna in whole-tree chambers under elevated atmospheric CO2 and summer drought[J].Plant,Cell and Environment,2012,35(5):966-981.
[2] 刘建锋,杨文娟,江泽平.等.遮荫对濒危植物崖柏光合作用和叶绿素荧光参数的影响[J].生态学报,2011,31(20):5999-6004.
[3] Senevirathna A M W K,Stirling C M,Rodrigo V H L.Growth,photosynthetic performance and shade adaptation of rubber (Hevea brasiliensis) grown in natural shade[J].Tree Physiology,2003,23(10):705-712.
[4] 李东胜,白庆红,李永杰,等.光照条件对蒙古栎幼苗生长特性和光合特征的影响[J].生态学杂志,2017,36(10):2744-2750.
[5] 贺顺钦,王发其.辽东栎苗木早期生长与光的关系[J].林业科学研究,2001,14(6):697-700.
[6] 陈秋夏,廖亮,郑坚,等.光照强度对青冈栎容器苗生长和生理特征的影响[J].林业科学,2011,47(12):53-58.
[7] 徐飞,郭卫华,徐伟红,等.不同光环境对麻栎和刺槐幼苗生长和光合特征的影响[J].生态学报,2010,30(12):3098-3107.
[8] Haller J M.Quercus virginiana:the southern live oak[J].Arbor Age,1992,12(5):30.
[9] Harms W R.Quercus virginiana Mill.live oak[M]// Burns R M,Honkala B H.Silvics of North America:Vol.2 hardwoods.Agric.handb.654.Washington,DC:U.S.Department of Agriculture,Forest Service,1990:751-754.
[10] 陈益泰,陈雨春,黄一青,等.抗风耐盐常绿树种弗吉尼亚栎引种初步研究[J].林业科学研究,2007,20(4):542-546.
[11] 陈益泰,孙海菁,王树凤,等.5种北美栎树在我国长三角地区的引种生长表现[J].林业科学研究,2013,26(3):344-351.
[12] Cavender-Bares J M,Kitajima K,Bazzaz F A.Multiple trait associations in relation to habitat differentiation among 17 oak species in North Central Florida[J].Ecological Monographs,2004,74(4):635-662.
[13] Spector T,Putz F E.Crown retreat of open-grown Southern live oaks (Quercus virginiana) due to canopy encroachment in Florida,USA[J].Forest Ecology and Management,2006,228(1-3):168-176.
[14] 王树凤,胡韵雪,李志兰,等.盐胁迫对弗吉尼亚栎生长及矿质离子吸收、运输和分配的影响[J].生态学报,2010,30(17):4609-4616.
[15] 王树凤,胡韵雪,孙海菁,等.盐胁迫对2种栎树苗期生长和根系生长发育的影响[J].生态学报,2014,34(4):1021-1029.
[16] 杨舒婷,曲博,李谦盛,等.弗吉尼亚栎幼苗对高温胁迫的生理响应[J].江西农业大学学报,2015,37(1):90-95.
[17] 宋以刚,邓敏,王妍婷.弗吉尼亚栎种子萌发特性[J].生态学杂志,2015,4(5):1295-1300.
[18] Ghasemi M,Arzani K,Yadollahi A,et al.Estimate of leaf chlorophyll and nitrogen content in asian pear (Pyrus serotina Rehd.) by CCM-200[J].Notulae Scientia Biologicae,2011,3(1):91-94.
[19] 叶子飘,于强.一个光合作用光响应新模型与传统模型的比较[J].沈阳农业大学学报,2007,38(6):771-775.
[20] 叶子飘,于强.光合作用光响应模型的比较[J].植物生态学报,2008,32(6):1356-1361.
[21] Maxwell K,Johnson G N.Chlorophyll fluorescence—A practical guide[J].Journal of Experimental Botany,2000,51(345):659-668.
[22] 张守仁.叶绿素荧光动力学参数的意义及讨论[J].植物学通报,1999,16(4):444-448.
[23] 吴飞燕,伊力塔,李修鹏,等.不同光照强度对石栎幼苗叶绿素含量及叶绿素荧光参数的影响[J].东北农业大学学报,2012,43(4):88-92.
[24] 潘瑞炽.植物生理学[M].北京:高等教育出版社,2008:61-62.
[25] 石玉波,卓丽环,辛雅芬,等.不同遮光处理对白栎光合生理特性的影响[J].安徽农业科学,2010,38(2):956-958.
[26] Dai Y J,Shen Z G,Liu Y,et al.Effects of shade treatments on the photosynthetic capacity,chlorophyll fluorescence,and chlorophyll content of Tetrastigma hemsleyanum Diels et Gilg[J].Environmental and Experimental Botany,2009,65(2-3):177-182.
[27] Falster D S,Westoby M.Tradeoffs between height growth rate,stem persistence and maximum height among plant species in a post-fire succession[J].Oikos,2005,111(1):57-66.
[28] Gardiner E S,Hodges J D.Growth and biomass distribution of cherrybark oak (Quercus pagoda Raf) seedlings as influenced by light availability[J].Forest Ecology and Management,1998,108(1-2):127-134.
[29] 闫兴富,曹敏.林窗对热带雨林冠层树种绒毛番龙眼幼苗生长的影响[J].应用生态学报,2008,19( 2):238-244.
[30] 杨兴洪,邹琦,赵世杰.遮荫和全光下生长的棉花光合作用和叶绿素荧光特征[J].植物生态学报,2005,29 (1):8-15
[31] 薛思雷,王庆成,孙欣欣,等.遮荫对水曲柳和蒙古栎光合、生长和生物量分配的影响[J].植物研究,2012,32(3):354-359.
[32] 郭志华,张旭东,黄玲玲,等.落叶阔叶树种蒙古栎(Quercus mongolica)对林缘不同光环境光能和水分的利用[J].生态学报,2006,26(4):1047-1056.
[33] Richardson A D,Berlyn G P.Spectral reflectance and photosynthetic properties of Betula papyrifera (Betulaceae) leaves along an elevational gradient on Mt.Mansfield,Vermont,USA[J].American Journal of Botany,2005,89(1):88-94.
[34] 梁文斌,聂东伶,吴思政,等.遮荫对短梗大参苗木光合作用及生长的影响[J].生态学杂志,2015,34(2):413-419.
[35] 唐敏,翟秀明,姚永红,等.不同茶树品种(系)叶绿素荧光参数季节性差异研究[J].西南农业学报,2015,28(1):79-83
[36] Yang H L,Yang X,Zhang Y G,et al.Chlorophyll fluorescence tracks seasonal variations of photosynthesis from leaf to canopy in a temperate forest[J].Global Change Biology,2017,23(7):2874-2886.
[37] Crous K Y,Zaragoza-Castells J,Ellsworth D S,et al.Light inhibition of leaf respiration in field-grown Eucalyptus saligna in whole-tree chambers under elevated atmospheric CO2 and summer drought[J].Plant,Cell and Environment,2012,35(5):966-981.